Aleshin V V
A.N. Belozersky Institute of Physicochemical Biology, M.V. Lomonosov Moscow State University 119992, Moscow.
Zh Obshch Biol. 2004 Jan-Feb;65(1):74-80.
The early embryonic development of Nematoda proceeds by three ways, which strictly correspond to three phylogenetic lineages. Under the first way the endodermal precursor is localized in the posterior blastomere at the two-cells stage (such a determination is the peculiarity of all the Chromadoria, including Secernentea and Caenorhabditis elegans). Under the second way the endodermal precursor is localized in the anterior blastomere of the egg. This feature is very unusual for Metazoa, but it is the only way of entoderm determination in all the Dorylaimia orders (Mononchida, Mermithida, Trichinellida, Dioctophymida, Dorylaimida). The third way described for the sea Enoplida is characterized with variable location of blastomers and changeable localization of endodermal precursor before eight-cells stage. It is still unknown of these three variants was typical the most recent common ancestor of present Nematoda. D.A. Voronov (2001) produced argument in favour of variable cleavage as primitive one for Nematoda. This opinion is rejected because of the similarity in development between sea Enoplida and C. elegans. Both of them share such features as low-cell gastrula and neurula, identical phylotypic lima bean stage of embryogenesis, identity of some geometrical figures 4 or 8 blastomers, isolating of the endodermal precursor at the eight-cells stage, the lack in development of any plesiomorphous features, which are widely distributed outside Nematoda (under the variable cleavage of Enoplida there are no such locations of blastomers, which are typical for spiral or radial cleavage, there are no embryonic leaves as well). One can see the homology of separate cells at adult Enoplida and Rhabditia. Cell lineage of Triplonchida as far as it is described at Tobrilus gracilis doesn't exclude the hypothesis on their origin from the cleavage similar to one of present Dorylaimia with localization of the endodermal precursor in the anterior blastomere. In view of all the considerations mentioned above one should interpret variable cleavage of Enoplida as derivation from invariant cleavage.
线虫的早期胚胎发育通过三种方式进行,这三种方式严格对应于三个系统发育谱系。在第一种方式下,内胚层前体在二细胞期定位于后裂球(这种决定是所有色矛纲动物的特点,包括尾感器纲和秀丽隐杆线虫)。在第二种方式下,内胚层前体定位于卵的前裂球。这一特征对于后生动物来说非常不寻常,但它是所有矛线目(单齿线虫目、索线虫目、旋毛虫目、膨结线虫目、矛线虫目)内胚层决定的唯一方式。描述的海线虫目的第三种方式的特征是,在八细胞期之前裂球位置可变且内胚层前体定位可变化。目前尚不清楚这三种变体中哪一种是现存线虫最近的共同祖先所特有的。D.A. 沃罗诺夫(2001年)提出论据支持可变卵裂是线虫的原始卵裂方式。但由于海线虫目和秀丽隐杆线虫在发育上的相似性,这一观点被否定。它们都具有低细胞原肠胚和神经胚、胚胎发育相同的系统型菜豆期、一些几何图形的4个或8个裂球相同、八细胞期内胚层前体分离、缺乏在其他线虫中广泛分布的任何近祖特征(在海线虫目的可变卵裂中,不存在典型的螺旋或辐射卵裂的裂球位置,也没有胚叶)。可以看出成年海线虫目和小杆线虫目的个别细胞具有同源性。就细弱托布线虫所描述的三齿线虫目的细胞谱系而言,并不排除它们起源于与现存矛线目类似的卵裂方式且内胚层前体定位于前裂球的假说。鉴于上述所有考虑因素,应该将海线虫目的可变卵裂解释为从不变卵裂衍生而来。